Poleward range expansion without a southern contraction in the

Transcription

ZooKeys 100: 333–352 (2011)
Poleward range expansion without a southern contraction
doi: 10.3897/zookeys.100.1535
www.zookeys.org
A peer-reviewed open-access journal
Research article
333
Launched to accelerate biodiversity research
Poleward range expansion without a southern
contraction in the ground beetle
Agonum viridicupreum (Coleoptera, Carabidae)
Claudia Drees1,6, Pietro Brandmayr2, Jörn Buse3, Petra Dieker4, Stephan Gürlich5,
Jan Habel4.6, Ingmar Harry7, Werner Härdtle6, Andrea Matern6, Hartmut Meyer6,
Roberto Pizzolotto2, Markus Quante6,8, Katharina Schäfer6, Andreas Schuldt6,
Angela Taboada9, Thorsten Assmann6
1 Tel Aviv University, George S. Wise Faculty of Life Sciences, Department of Zoology, The National Collections of
Natural History, Tel Aviv 69978, Israel 2 Università della Calabria, Dipartimento di Ecologia, I-87036 Arcavacata di Rende (CS), Italy 3 Johannes Gutenberg-University Mainz, Institute of Zoology, Department of Ecology,
Becherweg 13, D-55099 Mainz, Germany 4 Musée National d’Histoire Naturelle, Section Zoologie des Invertébrés, L-2160 Luxembourg, Luxembourg 5 Verein für Naturwissenschaftliche Heimatforschung zu Hamburg
e.V., Martin-Luther-King-Platz 3, D-20146 Hamburg, Germany 6 Leuphana University Lüneburg, Institute of
Ecology and Environmental Chemistry, Scharnhorststr. 1, D-21335 Lüneburg, Germany 7 ABL, Nägeleseestraße
8, D-79102 Freiburg, Germany 8 GKSS Research Center, Institute for Coastal Research, Max-Planck-Straße 1,
D-21502 Geesthacht, Germany 9 Area of Ecology, Department of Biodiversity and Environmental Management,
University of León, Campus de Vegazana s/n, E-24071 León, Spain
Corresponding authors: Claudia Drees ([email protected]), Thorsten Assmann ([email protected])
Academic editor: R.Vermeulen | Received 4 January 2010 | Accepted 23 March 2010 | Published 20 May 2011
Citation: Drees C, Brandmayr P, Buse J, Dieker P, Gürlich S, Habel J, Harry I, Härdtle W, Matern A, Meyer H, Pizzolotto
R, Quante M, Schäfer K, Schuldt A, Taboada A, Assmann T (2011) Poleward range expansion without a southern
contraction in the ground beetle Agonum viridicupreum (Coleoptera, Carabidae). In: Kotze DJ, Assmann T, Noordijk J,
Turin H, Vermeulen R (Eds) Carabid Beetles as Bioindicators: Biogeographical, Ecological and Environmental Studies.
ZooKeys 100: 333–352. doi: 10.3897/zookeys.100.1535
Abstract
We investigated the extent of poleward shifts in the distribution range of Agonum viridicupreum due to climate change in the western Palaearctic. Species’ records were obtained from extensive literature sources as
well as from collections, and consistent amateur entomologists’ recordings. Within the general geographic
range of the species, we analyzed in detail two parts of both, the northern and southern distribution range
boundaries: (1 and 2) north-western Germany (leading or high-latitude edge), (3) Israel and (4) southern Italy (rear or low-latitude edge). Temporal changes in the occurrence data of the species indicated a
northward shift of the leading edge of a minimum of 100 km within the last 50 to 100 years. In contrast,
according to the data gathered, the rear edge has not changed during the last decades. Further studies are
Copyright Claudia Drees et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which
permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
334
Claudia Drees et al. / ZooKeys 100: 333–352 (2011)
needed in order to fully understand the underlying mechanisms of the different behaviour of leading and
rear range edges of A. viridicupreum in the current context of global change. Despite our incomplete understanding, chronosequences of the occurrence of the given species have the potential to optimize climate
niche modelling to predict trends in the distribution range in the future.
Keywords
chronosequence, climate change, distribution area, global change, wetlands, power of dispersal, migration, range shift
Introduction
For about 250 years, man has released radiatively active gases and particles in substantial amounts into the atmosphere. As one of the consequences, the global mean near
surface temperature has increased, a phenomenon commonly referred to as ‘global
warming’ or ‘climate change’. Deduced mainly from instrumental observations initiated around 1860, the observed climate change can be attributed to a large extent to
human activities, which influence not only global temperature, but also pH-values of
the oceans, precipitation and the general hydrological cycles on Earth (IPCC 2007;
Quante 2010).
For many animal and plant species, theoretical analyses on the climate determination of the species’ occurrence have predicted a general poleward shift and (in mountain
areas) an uphill shift of the given distribution areas and populations, respectively, as a
response to climate change. In agreement with theory, numerous range shifts have been
documented in the last years. Examples are known from vascular plants, birds, and many
insects such as butterflies, dragonflies and damselflies (Hickling et al. 2006; Parmesan
2006; Pauli et al. 2007). Carabid beetles and other epigean soil invertebrates are well
known as highly dynamic colonizers of glacier forelands in the last two centuries, and
uphill shifts of several hundred metres altitude have been described in the Austrian and
Italian Alps and for Scandinavian mountains (Gobbi et al. 2006; Gobbi et al. 2007).
Poleward shifts of distribution areas are very likely also for widely distributed carabid species (in contrast to species with restricted distribution areas, i.e. endemics), as
their patterns of geographic distribution are strongly determined by climatic factors (as
shown by a large-scale analysis of West Palaearctic ground beetle diversity, Schuldt and
Assmann 2009). Indeed, northward shifts of ground beetle species have been documented several times in the literature. Already Lindroth (1972), certainly the most
important carabid biogeographer, demonstrated that several species, especially those
with flight activity, have expanded their distribution areas northwards in Fennoscandia
since the middle of the last century. Some of these species went on spreading polewards, e.g. Stenolophus mixtus in Scandinavia (Kvamme 1978; Palm 1982) or in Great
Britain (Blake 2001).
Moreover, ground beetles with their northern distribution limit in Britain have
moved about 50 km northwards within a period of about two decades (Hickling et al.
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2006). Further examples of poleward shifts in the geographic distribution of carabids
can be obtained from the faunistic literature throughout Europe, e.g. Demetrias imperialis in countries around the Baltic Sea (Silfverberg 2005), and Tachyta nana, Diachromus germanus, and Acupalpus luteatus in north-western German lowlands (Ziegler
2004). Besides, expansions of carabid species’ distribution areas are conspicuous and
numerous amateur entomologists consistently notify new records.
However, previous studies on poleward range margin shifts of ground beetles have
mainly focused on the leading (i.e. current high altitudinal and latitudinal) edges of
their distribution areas (literature cited above). Changes occurring at the leading edge
are interesting, especially in the framework of dispersal biology, and they enable us to
understand many population biological processes (Hengeveld 1985; 1989). In contrast, despite the fact that leading edges seem to be more relevant than rear (i.e. current
low altitudinal and latitudinal) edges, the latter may be of greater importance for the
long-term survival of species (Hampe and Petit 2005). This is related to the different
histories of leading and rear edges. In general, at the poleward limits of distributions
newly founded populations are recent and, therefore, only short-term adaptations have
been possible. In contrast, many of the rear edge populations are close to their glacial
refuges, i.e. the specimens are genetically more variable and, thus, allow greater power
of adaptability and preadaptation (Hampe and Petit 2005).
Nevertheless, up to now, there is no available study comparing the reaction of a
ground beetle species at both margins of its distribution range. Thus, in this study we
aimed at investigating the extent of poleward range shifts at both the leading and rear
edges of the distribution area of a carabid species due to recent climate changes. We
selected Agonum viridicupreum as our study object because it fulfils many preconditions of a suitable model species to assess potential margin shifts: The specimens can be
easily found in the field, they are fully winged and fly actively, and the species’ habitat
preferences are well-known. Furthermore, the specimens are nicely coloured, stimulating many amateur entomologists to record the species, and, therefore, allowing suitable faunistic data from large parts of its distribution area. Moreover, the species is not
restricted to habitats that are influenced or even destroyed by other drivers of global
change, nor have been altered simultaneously by the temperature increase in the last
decades (e.g. oligotrophic peat bogs affected by increased atmospheric nitrogen depositions due to pollution, Bobbink et al. 1998).
Material and methods
The study species
Agonum viridicupreum (Goeze, 1777) is a macropterous and thermophilous species
restricted to open, wet habitats such as meadows, fens and rain ponds. The day-active
beetle prefers sun-exposed muddy sites where it can be easily detected by its greenbronze-coloured surface. Due to its occurrence in floodplain areas (with high prob-
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ability of diversion), the dispersal of individuals is not only determined by the species’
ability to fly. Specimens can be transported downstream by flooding events into areas
where the species might not be able to establish autochthonous populations (Bonn
2000; Turin 2000; personal observations).
In the Levant (Middle East, see below), the beetle lives in wet habitats, mostly close
to winter or rain ponds (personal observations). In southern Italy (Calabria) the species
lives in river bank habitats around Typha swamps or in other wet vegetation types and
crops, and on lake shores, until about 1400 m above sea level.
Distribution area and temporal changes
We reviewed the available faunistic literature for the western Palaearctic (Europe,
the Mediterranean area) to determine the general distribution area of the study species (Horion 1941; Jeannel 1941f; Antoine 1955f; Kocher 1963; Magistretti 1965;
Bonadona 1971; Burakowski et al. 1973f; Alfieri 1976; Bangsholt 1983; Lindroth
1985; Jeanne and Zaballos 1986; Hieke and Wrase 1988; Marggi 1992; Zaballos
and Jeanne 1994; Guérguiev and Guérguiev 1995; Kryzhanovskij et al. 1995; Hurka 1996; Machard 1997; Köhler and Klausnitzer 1998; Casale and Vigna-Taglianti
1999; Drovenik and Peks 1999; Neculiseanu and Matalin 2000; Turin 2000; Marggi
and Luka 2001; Bousquet 2003; Serrano 2003; Müller-Motzfeld 2004; Brandmayr
et al. 2005; Curcic et al. 2007; Luff 2007; Austin et al. 2008; Desender et al. 2008)
and of Agonum fulgidicolle Erichson, 1841, an allopatric sibling taxon of A. viridicupreum (ranked by some authors as a subspecies, e.g. Puel 1938), which occurs in
north-western Africa.
The situation of faunistic recordings is sufficient for one region at the northern
distribution edge (north-western Germany) and for two regions at the southern distribution edge (Levant in the Middle East, mainly Israel, and Calabria in southern Italy).
– North-western Germany has been studied by numerous amateur entomologists
who have greatly contributed to our knowledge on the geographic distribution
of carabid beetles. We therefore analyzed the changes in the species’ distribution
separately for (a) West Lower Saxony (west of river Weser) and for (b) East Lower
Saxony (east of river Weser), Hamburg, and Schleswig Holstein. For these regions
records from three periods (before 1950, between 1951 and 1980, after 1980) were
summed up to document tendencies in the numbers of catches.
– For Israel, the first records date from the 1920s (the beginning of modern zoological exploration of the given region by local scientists, in former times only
explorers from abroad collected beetles there). We therefore distinguished only two
periods of collecting: before 1980 and after 1980.
– For southern Italy (Calabria) there are scarce historical records (before 1980).
However, after 1980, intensive ecological surveys were carried out on populations
in several sites of the Crati river valley (Mazzei et al. 2010).
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Consequently, within the global distribution area of the study species, we analyzed the
northern and southern range boundaries by studying in detail the three above mentioned
concrete margin regions, where the coverage of the faunistic recordings is amply and sufficient: (1 and 2) a part of the leading edge (north-western Germany: Lower Saxony, Hamburg and Schleswig-Holstein, divided into regions west and east of the river Weser), and
(3 and 4) the only continental areas at the rear edge that are not limited by the sea or by
the presence of A. fulgidicolle (the Levant in the Middle East: mainly Israel, and Calabria,
(southern Italy)). For these areas we compiled numerous faunistic records mostly published in local journals (Westhoff 1881, 1882; Bodenheimer 1937; Barner 1954; Lohse
1954; Gersdorf and Kuntze 1957; Assmann and Ehrnsberger 1990; Assmann and Ehrnsberger 1990; Angelini 1991; Assmann 1991; Mossakowski 1991; Gürlich et al. 1995;
Handke 1995; Handke and Kundel 1996; Bonn et al. 1997; Fuellhaas 1997; Nitzu 1997;
Ziegler 1997; Fischer et al. 1998; Bonn 2000; Hannig and Schwerk 2000; Hannig 2001;
Bonn et al. 2002; Assmann et al. 2003; Hannig 2004; Günther and Assmann 2005;
Hannig 2005, 2008; Wrase 2009; Mazzei et al. 2010). At the leading edge, Assmann and
Ehrnsberger (1990) as well as Irmler and Gürlich (2004) have previously observed an
enlargement of the distribution range in northern Germany. Moreover, we also incorporated in our data base the species’ records obtained from museums and private collections
(collections of several authors and David Wrase, Berlin (CWB), the Collection Assmann
Bleckede (CAB), The National Collection of Natural History of the Tel Aviv University
(TAU)) and data bases available on the internet (mainly www.entomologie.de/hamburg/
karten/%0bfhl_02/_agovir1.htm and www.eurocarabidae.de). Generally, several specimens from identical dates and locality are regarded only as one record.
Climate changes in the regions of interest
We surveyed climatological literature and compiled information about recent climate changes in the three regions north-western Germany, Israel and southern Italy
(Calabria). We focussed only on changes in temperature and precipitation, the main
factors influencing the ground beetles’ biology and distribution.
Results
Climate changes
North-West Germany – warmer springs with wetter winters and drier summers
Over the last 150 years a considerable increase of the global mean temperature by about
0.8°C has been observed. Also for western Europe the measurements show a warming
trend. For Germany during the 20th century a mean temperature rise of about 1.0°C
338
was reported by Schönwiese and Janoschitz (2008). This warming is not homogenous;
there are noticeable seasonal and regional differences. In the western part of northern
Germany a linear trend value for the temperature between 0.6°C and 0.8°C appears to
be typical. For the period from 1951 to 2000 this linear trend value is slightly higher
and comes close a 1°C with a tendency of marginally higher values towards the southeast. The increase in winter temperatures was higher than that for the summer. For the
last decades the strongest warming was found to appear in spring. An evaluation of station data for different states in northern Germany using a different averaging method
came to the conclusion that during the 20th century the mean temperature in Lower
Saxony rose by 1°C, in Schleswig Holstein by 0.8°C and in the metropolitan region of
Hamburg by 1.1°C (I. Meinke, GKSS pers. comm.). For the Hamburg area Schlünzen
et al. (2010) report an increase in the decadal warming rate, which underlines that the
temperature trend was significantly larger in the last three decades. The corresponding
rates from a piecewise linear trend evaluation are 0.07 K/decade for 1891–2007, 0.19
K/decade for 1948–2007 and 0.60 K/decade for 1978–2007. Recently the strongest
warming appeared in the winter months. A comparison of mean temperatures for the
first and last decade of the 20th century suggests that the region in Lower Saxony west
of the river Weser faced a slightly higher warming than the eastern part. This result is
in conflict with the pattern shown by Schönwiese and Janoschitz (2008) and probably
due to the method of comparing only two decades.
Linear 20th century precipitation trends for Germany have been reported to be
about 8.5% (an increase from 750 mm to 800 mm, Schönwiese and Janoschitz 2008).
However, because of a strong interannual variability this trend is not statistically significant. Over this period especially the winter precipitation increased, while for the summer months a decrease was observed. This increase in winter precipitation and decrease
in summer precipitation was also reported for the western part of northern Germany.
An evaluation of station data for different states in northern Germany using a different
averaging method came to the conclusion that during the 20th century precipitation in
Lower Saxony increased by about 10%, in Schleswig Holstein by about 12.5% and in
the metropolitan region of Hamburg by about 12% (I. Meinke, GKSS pers. comm.).
For the Hamburg area Schlünzen et al. (2010) report a significant increase in precipitation rate. The corresponding rates from a piecewise linear trend evaluation are ~0.8
mm/year for 1891–2007 and 1.3 mm/year for 1948–2007. The increase again is most
pronounced for the winter months. For the months April and July in the period between
1978 and 2007 a significant decrease in precipitation in the Hamburg area has been
found.
Levant (Israel): warmer and drier in the north, wetter in the south
An analysis for the period 1964 to 1994 of temperature measurements at 40 stations
evenly distributed over Israel came to the conclusion that there appears to be a general
warming trend, with some local exceptions, i.e. in the south, which could be related
339
to enhanced aerosol emission (Ben-Gai et al. 1999). This general trend has been confirmed by a more recent reanalysis study (Saaroni et al. 2003); this study also notes that
for the last decades July replaces August as the warmest month of the year. The overall
analysis reveals a complex change pattern. First, the summers have become warmer,
while the winters became colder; second, there exists a significant decreasing trend of
the daily maximum and minimum temperature during the cool season and an increasing trend during the warm season (Ben-Gai et al. 1999).
Concerning climatological precipitation trends the Levant has to be divided into
a southern and northern part. An analysis of winter half-year precipitation over the
entire Mediterranean region reveals predominating rainfall decreases during the last 50
years. The areas deviating from this general trend includes southern Israel (Jacobeit et al.
2007). Several studies report opposing trends of annual rainfall for the eastern Mediterranean (e.g. Steinberger and Gazit-Yaari 1996; Jacobeit et al. 2007; Khatib et al. 2007),
a decrease of rainfall amounts in the northern part of Israel and increase for southern
regions during recent decades. There are indications that the observed trend differences
are the outcome of changes in synoptic conditions in the eastern Mediterranean region
(Steinberger and Gazit-Yaari 1996). In the overall series of wettest winters (see above,
analysis by Luterbacher et al. 2006) the southern part of the Levant was slightly drier
than the climatological mean and in the overall driest winter series this region was wetter than the 1961 to 1990 average (Luterbacher et al. 2006). For the northern part of
the Levant the trends seem to be vice versa; consistently different trend behaviour in the
southern part compared to the northern part of Levant has been observed.
Calabria: Warmer and drier
From the maps of linear trends in annual mean temperature for Europe compiled
by Schönwiese and Janoschitz (2008) for Calabria a warming trend of about 1°C for
the entire last century can by extracted, the value is consistent with the analysis by
Gerstengarbe and Werner (2007), who compared the first and last decades of the 20th
century. The respective value for the period from 1951 to 2000 is slightly larger than
0.6°C. This annual mean temperature trend does not reflect seasonality; warming was
driven mainly by the summer months while for the winter months even a slight cooling trend was observed.
Overall it can be said that the most southern part of Italy and especially Calabria
has become drier over the last decades. While the linear trend in annual precipitation
for the entire 20th century for the Calabria region is almost zero, a pronounced trend
exists for the period from 1951 to 2000 with a decrease in precipitation by about 20%
in the annual mean with a decrease in summer precipitation of about 40% (Schönwiese and Janoschitz 2008). A comparison of the first three decades with the last three
decades of the 20th century reveals a slightly drier Calabria at the end of the century
(Gerstengarbe and Werner 2007). Luterbacher et al. (2006) analyzed winter precipitation anomalies for the last centuries in the Mediterranean region. The wettest decade
340
was 1961 to 1970 and the driest was 1986 to 1995. The wettest (driest) multidecadal
periods (30 winters in a row) were from 1951 to 1980 (1973 to 2002) with 5 mm (-15
mm) departures from the climatological average (1960 to 1990). Interestingly, in the
overall wettest winters Calabria was drier than the climatological mean (10 to 20 mm)
and in the overall driest winter series Calabria was about 10 to 20 mm wetter than the
61 to 90 average.
Geographic and altitudinal distribution area of A. viridicupreum
The distribution area of the species within the western part of the Palaearctic is given
in Fig. 1. A. viridicupreum occurs around the Mediterranean Sea (with a distribution
gap in north-eastern Africa). The northern edge of the distribution area runs from the
Netherlands through northern Germany and Poland (see also Fig. 2). In the south-east
the species occurs in Turkey, Lebanon, and Israel.
In the southern Iberian Peninsula and Morocco, the beetle prefers mountainous
areas (Zaballos and Jeanne 1994), but in the central and northern parts of Spain it also
thrives well in lowland habitats (down to sea level, e.g. close to Oviedo, in the mountains up to about 2000 m a.s.l., CAB). In south-eastern Europe, the species occurs in
mountains as well as lowlands (e.g. Peloponnese, CAB).
The south-eastern distribution edge in the Levant virtually coincides with the border of the Mediterranean climate (Fig. 3). In this study, we report the first record for
Egypt ([(T)El Arish, Sinai, leg. L. Fishelsohn, 12.03.1956], record in TAU, Fig. 3).
However, the single specimen collected is not a proof of the existence of an autochthonous population here. The same is true for records obtained from the desert regions
(e.g. Dead Sea Region, where no suitable habitats for the species occur, cf. Fig. 3).
Faunistic analyses of the distribution margins
West Lower Saxony (west of the river Weser): Although A. viridicupreum has been
known from the Netherlands since the 19th century, no specimens were recorded from
West Lower Saxony until the 1980s (Table 1). Indeed, in the 1950s the northern
distribution limit of the species’ range was located southwards of Lower Saxony, in
the Westphalian Lowlands (Horion 1941; Barner 1954; Westhoff 1881). However,
after 1981 numerous records from the whole Lower Saxonian (and Westphalian) Lowlands, northwards to the North Sea, were reported (Assmann and Ehrnsberger 1990;
Mossakowski 1991; Handke and Kundel 1996; Fuellhaas 1998; Günther and Assmann 2005; Hannig and Schwerk 2000; Hannig 2001, 2005, 2008, numerous records
in collections, e.g. CAB), thus, expanding the former northern distribution margin
(Fig. 2). The distance between the known northern limit of the 19th century (central
Westphalian Lowlands) and the present records close to the North Sea coast is more
than 100 km (Fig. 2).
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Figure 1. Distribution of Agonum viridicupreum (shaded in grey) and its sister taxon A. fulgidicolle in
the western Palaearctic. Map modified after Turin et al. (2003) using information from Brandmayr et al.
(2005) and personal observations. Frames indicate regions selected for more detailed analyses of records,
see Figs 2 and 3.
East Lower Saxony (east of river Weser): Horion (1941) and Gersdorf and
Kuntze (1957) listed records of the species from the hilly countryside close to Hannover. The latter authors questioned the occurrence of the species in the lowlands of
eastern Lower Saxony. Along the river Elbe, one old record (19th century) is known
from one site south-east of Hamburg (“?” in Fig. 2, Table 1). Lohse (1954) interpreted
the presence of these specimens as vagrants transported downstream by flooding events
from south-eastern Germany. However, these specimens could have originated also
from temporal populations.
Between 1951 and 1980, only one record from another site in the Lower Saxonian
floodplain area of the river Elbe is known (Table 1). Records from sites outside the
given floodplain are exclusively known since 1981, when the number of records greatly
increased.
Today, the species is found northwards, up to central Schleswig-Holstein (www.
entomologie.de/hamburg/karten/fhl_02/_agovir1.htm), and reaches also the northwestern parts of the considered area. Interpreting the old records from the floodplain
area of the river Elbe as autochthonous populations leads us to think that the species’
geographic range has experienced a northward shift of about 100 km during the last
century. Even if these records were not seen as autochthonous populations, the shift
would have spanned over about 200 km (Fig. 2).
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Figure 2. Distribution of A. viridicupreum (shaded in grey) in North-West Germany with eastern parts
of the Netherlands. Arrows indicate minimum range expansion in the last three decades (for explanation
and records see text). Range expansion in the Netherlands indicated after Turin, pers. comm.
Levant: Bodenheimer (1937) listed A. viridicupreum from Israel for the first time, and
the former documented records were taken in the 1920s (TAU). In this region, the beetle is
abundant at many rain or winter ponds (up to ca. 20 individuals per hour can be collected
by hand picking; personal observations). So, it is very likely that the late discovery of the
species at its south-eastern distribution edge would be a consequence of the poor carabidological exploration of the country. Since Bodenheimer’s time, numerous new records of the
species have been reported, also during the last years (Fig. 3). A. viridicupreum reaches the
south-eastern limit of the Mediterranean climate in Israel. There is no evidence for a northward shift of its distribution range, as the known southern Israeli populations are close to
the semi-arid climate region from where the species is virtually unknown; only singletons
– not indicating autochthonous populations – have been found (see above and Fig. 3).
Calabria: For southern Italy (Calabria) there are scarce historical records (before
1980; Magistretti 1965; Angelini 1991). After 1980, a total of 37 specimens were recorded (Mazzei et al. 2010).
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Figure 3. Distribution of A. viridicupreum in Israel. The striped area indicates Mediterranean climate
zone (according to Yom-Tov and Tchernov 1988). Species’ records are taken from collections TAU, CAB
and CWB.
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Table 1. Number of records of A. viridicupreum in the different periods of time in north-western
Germany (leading edge) and Israel (rear edge).
Study region
Leading edge:
North-West Germany – West Lower Saxony
North-West Germany – East Lower Saxony
and Schleswig Holstein
Rear edge:
Israel
Number of records
before 1950 1950–1980
after 1980
0
11
0
12
12
26
24
14
Close to Geesthacht (leg. Kolze, 1890 [river Elbe, east of Hamburg], because of lacking records from the
surrounding seen as diversion by Lohse, 1954)
2
Pevestorf [river Elbe, south-east of Lüneburg]
1
Discussion
Poleward shift of the leading edge
The compilation of the faunistic data showed that the distribution range of A. viridicupreum had significantly shifted northwards within the last 50 to 100 years. Up
to 1950, in the analyzed region, the northern edge of the species’ distribution had
stretched from the Netherlands (Nijmegen, Enschede; Turin 2000) southwards to the
Westphalian Lowlands (south-western Lower Saxony), and again northwards to Hannover and Braunschweig (Fig. 1). The historical lack of the species studied in the region
around Osnabrück (south-western Lower Saxony) does not need to be the result of
undersampling, as also other thermophilous insects colonized this region later than the
neighbouring western or eastern regions (e.g. several grasshopper species; Hochkirch
2001; Hochkirch and Damerau 2009).
Today, in the western part of Lower Saxony, A. viridicupreum can be found up
to the North Sea, confirming a northwards range expansion of about 100 km. Similarly, in the neighbouring Netherlands the beetle has expanded its distribution range
northwards and can nowadays be recorded close to the city of Groningen (Turin, pers.
comm.). Also in the eastern part of Lower Saxony the species has spread northwards a
minimum distance of 100 km and it reaches the centre of Schleswig-Holstein today.
These results allow the assumption that a further temperature rise will make the species’ occurrence in Denmark highly probable in the near future.
Stable rear edge
Unlike the northern distribution edge, the southern range margin (rear edge) of A. viridicupreum has not changed within the last decades. Indeed, there are still populations
with numerous individuals south of Tel Aviv, which is close to the southern limit of the
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Mediterranean climate. Consequently, in Israel we expect the beetle to occur in most
of the regions characterized by Mediterranean climate. In southern Italy (Calabria)
A. viridicupreum shows a stable rear edge north of the 39th parallel, with permanent
populations in the Crati Valley in the Cosenza province, in areas marked by Submediterranean or Mediterranean climate.
Different behaviour of leading and rear edges
In accordance with numerous other authors (e.g. Hengeveld 1985; Hickling et al. 2006),
we interpret the poleward shift of the leading edge of this species as a consequence of
increasing temperature. It seems to be more difficult to describe the differences between
leading and rear edge. In general, they could be explained by either intrinsic or extrinsic
factors, or even a combination of both. The influence of intrinsic factors would imply that
distinct genetic make-ups of the populations from the opposite edges of the distribution
range are likely (Hampe and Petit 2005). However, so far there are no available investigations to corroborate this hypothesis for A. viridicupreum or other ground beetle species
with the tendency of invasions in the Levant. Consequently, additional investigations are
necessary. For non-migratory butterflies, it has been demonstrated that population size
fluctuations are more pronounced at the leading edge than at the rear one (Parmesan et
al. 1999) – a possible indication of less well-adapted populations at the leading edge. As
our data do not give information about the population sizes of A. viridicupreum at the
various sites, on the one hand, we cannot investigate this assumption. On the other hand,
the high catching rates (which are comparable to catches from sites in northern Italy and
Germany) do not support the assumption of the species declining in the Levant.
On the contrary, extrinsic factors may be acting differently at the leading and rear
edge of the distribution range of A. viridicupreum. Our compilation of recent climate
trends, however, reveals generally rising temperatures in all regions under study. In
contrast, less consistent changes in precipitation can be observed. The populations
in both, Calabria and the Levant will have to deal with a reduction in mean annual
precipitation. In the face of the virtual exclusive occurrence of the species close to rain
ponds in the southern edge of its distribution, it seems likely that the southern populations are limited by the given ground water tables which predominantly result from the
annual amount of precipitation (mainly in the winter months).
In the Levant, larval development takes place during winter and early spring, as revealed
by numerous tenerals, even at higher altitudes, e.g. 900 m a.s.l., in the Golan Heights, in
April and May (personal observation). In contrast, the northern populations in Central
Europe are unlikely to be limited by the amount of precipitation, but rather by temperatures during the species’ activity period. In fact, in this region, larval development takes
place during summer and tenerals occur in late summer and autumn (August to October;
Turin 2000; personal observations; during this season tenerals has never been found in the
Levant). Finally, the role of other factors such as interactions with other organisms cannot
be excluded when interpreting the distribution changes at the species’ range margins.
346
Nonetheless it is possible that the northern and southern limits of Agonum viridicupreum are determined by different climatic factors: increasing summer temperature
in the north and increasing precipitation during the winter in the south can explain the
poleward shift of the leading and the stable rear edge of the given species’ distribution.
Potential of A. viridicupreum for further ecological research on global change
This study is the first one that investigates simultaneously the possible shifts of the
northern and southern margins of a carabid species’ distribution due to climate change.
Undoubtedly, at present we are not able to fully understand the underlying mechanisms of the different behaviours of the leading and rear boundaries of the geographic
range of A. viridicupreum in the actual context of global change. However, our analysis
suggests that the reaction of the study species to climate change may be more intricate
than expected at first. For this reason, we think that the more complex situation in A.
viridicupreum has important potential for further carabidological investigations at the
interface of global change ecology and conservation biology. For instance, predictions
based on climate envelope modelling, which has become both commonplace for many
other animal species and the object of an intensive (and critical) scientific discourse
(Settele et al. 2009; Rödder and Dambach 2010), can be optimized (and evaluated) by
using the chronosequences of distribution data. To our knowledge, this approach has
not yet been applied for ground beetles, although they appear to be an excellent object
to validate climate envelope models, thanks to the outstanding faunistic work with
numerous records from many regions and time periods (e.g. Luff 1998; Turin 2000;
Desender et al. 2008; Trautner, in prep.).
Acknowledgements
We thank Pascale Zumstein (Lüneburg) for her help with compiling records, the
Hamburg Coleopterists group for providing additional records and Ariel-Leib-Leonid
Friedman (Tel Aviv University) for helping with reading old Hebrew labels. Claudia
Drees is a VATAT funded post-doctoral fellow at the Tel Aviv University Zoological
Museum.
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Poleward range expansion without a southern contraction in the

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